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The Theropod Ancestry of Birds: New Evidence from the Late Cretaceous of Madagascar
C. A. Forster, S. D. Sampson, L. M. Chiappe, and D. W. Krause
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Supplemental Data
Character matrix for phylogenetic analysis of Rahona ostromi.
5 10 15 20 25 30 35 40 45
50 55 60 65 70 75 80 85 90
95 100 105 110 113
Allosaurus 00000 00000 00000 00000 00000 00000 00000 00000 00000
00000 00000 00000 00000 00000 00000 00000 00000 00000
00000 00000 00000 00000 000
Compsognathus 0100? 00000 10??? ?0?11 01??0 ?00?0 ??00? 00??0 ?0000
?0000 ???0? ?0000 00??0 0?00? 000?? 1?100 10?00 0??0?
????? ????? ?00?? 00000 1?0
Archaeopteryx 02111 00111 10110 01?11 01000 100?1 00101 1?101 10010
11010 01110 00010 01112 10111 11001 11111 12110 11000
111?? 0111? ?0111 00100 100
Rahona ????? ????? ????? ????? ????? ??111 11?01 10??1 10011
????0 ????? ??01? ????2 ??11? 11101 11111 12110 11000
21200 01112 11010 0?101 100
Oviraptoridae 1???1 10101 10110 00001 01001 00101 10000 00000 10000
10000 01100 00000 00001 00001 00000 00000 11000 00000
11000 00000 00000 00000 100
Dromaeosauridae 01001 10001 00110 11011 01000 00101 00001 10001 00000
10010 01010 00000 01111 00011 00000 10110 11000 01000
11100 00000 00010 00001 000
Ornithomimidae 02011 11010 10110 10111 01001 00000 00000 00010 00000
10000 00000 00000 00011 00000 00000 00010 11000 00000
11000 00000 00001 00010 000
Troodontidae 00011 01111 001?0 ?0111 11001 ?0100 10001 10011 00000
10000 00?10 00010 01011 10011 0000? ?0011 11000 01000
11200 00001 00110 0?011 100
Tyrannosauridae 00001 11101 00000 00000 10010 00001 00000 00000 00000
00000 00000 00000 00020 00000 00000 00100 10000 00000
00000 00000 00000 00010 000
Alvarezsauridae ?1111 01?10 11??1 11011 1?001 ?0110 11101 10110 11000
10000 00211 00001 0??20 11101 0??1? 02201 2??00 00211
21000 00111 11110 00?10 100
Enantiornithes 01111 01?10 111?? 1???1 ????1 ?1110 11211 ?111? 10111
21120 12211 11111 11112 00111 00111 0?201 2?101 11100
21?11 1011? 11211 01100 100
Ornithurae 11111 11110 11021 1?011 11111 11110 11211 ?111? 10101
21121 12211 11111 11112 01111 00211 02201 2?001 11211
21011 10112 11211 11100 111
Unenlagia ????? ????? ????? ????? ????? ???01 11??? ?0??? ??01?
????0 ????0 0???? ????? ????? 11001 11111 11100 11000
11000 01??? ????? ????? ???
Patagopteryx ????? ?1?10 ????1 1???1 ??11? ?1110 11??? ?111? 11111
21121 1?11? ??011 ??0?? 0111? 00010 02201 20001 10211
210?1 10011 11211 11000 111
Iberomesornis ????? ????? ????? ????? ????? ?01?0 0011? ???1? 101??
21?20 12??? 00?1? ????? ???1? 0???? ??1?? 2?10? 1????
?1??? ??1?? ????0 0?100 1?0
Character List for phylogenetic analysis of Rahona ostromi
dental
- teeth in premaxilla: present (0); absent (1).
- denticles on teeth: present on anterior and posterior carinae (0); present on posterior carina, but absent or severely restricted on anterior carina (1); absent (2). (adapted from Martin et al., 1980)
- shape of teeth: laterally compressed and recurved (0); slightly compressed, nearly conical (1). (Ostrom, 1976)
- constriction between crown to root: absent (0); present (1). (Martin et al., 1980)
skull
- jugal contribution to antorbital fenestra: absent, eliminated by lacrimal-maxilla contact (0), present (1). (Holtz, 1994)
- maxilla contribution to naris: absent, excluded by a nasal-premaxilla contact (0); present (1). (Martin et al., 1980; Currie, 1995)
- shape of frontals: anteriorly broad (0); triangular, narrow rostrally (1). (Currie, 1987b; Russell and Dong, 1993)
- extreme reduction or loss of prefrontal: absent (0); present (1). (Gauthier, 1986)
- frontal length relative to parietal: smaller or subequal (0); nearly two times as long (1).
- narrow sagittal crest on parietals: absent (0); present (1). (Currie and Zhao, 1993)
- jugal bar shape: robust (0); thin and straight (1). (Ostrom, 1976)
- jugular postorbital process: present (0); absent (1). (Chiappe)
- subsidiary foramen between palatine and pterygoid: absent (0); present (1). (Gauthier, 1986)
- palatine-ectopterygoid contact: no contact, ectopterygoid present (0); contact present, ectopterygoid present (1); no contact, ectopterygoid absent (2). (adapted from Currie, 1995)
- medial otic process of quadrate that articulates with prootic: absent (0); present (1). (Chiappe, 1995)
- distal quadrate articulation set well anterior to proximal quadrate cotyle: absent (0); present (1). (Ostrom, 1976)
- shape of paroccipital process: straight and vertically oriented (0); distal end with distinct twist as to face caudodorsally (1). (Colbert and Russell, 1969; Currie, 1995)
- bulbous parasphenoid capsule (cultriform process): absent (0); present (1). (Osmolska et al., 1972)
- accessory lacrimal fenestration: present (0); absent (1).
- supraoccipital crest: absent or weak (0); well pronounced (1).
- fusion between parietals and laterosphenoid in adults: present (0); absent (1).
- accesory fenestra between premaxilla and maxilla: present (0); absent (1).
- quadratojugal cotyla in the lateral face of the mandibular process of the quadrate: absent (0); present (1).
- pneumatic quadrate: absent (0); present (1).
lower jaw
- unfused interdental plates: present (0); absent (1). (Currie, 1987; Holtz, 1994)
- separate coronoid bone: present (0); absent (1). (Elzanowski and Wellnhofer, 1996)
vertebral column
- heterocoelus cervical vertebrae: absent (0); present (1). (Sanz et al., 1995)
- ventral processes (hypapophyses) on cervicodorsal vertebrae: absent or very weakly developed (0), present, well developed (1). (Gauthier, 1986)
- ratio of height of neural canal in dorsal vertebrae to height of cranial articular face: less than .40 (0); more than .40 (1). (Sanz et al., 1995)
- pneumatic foramina on dorsal vertebrae: absent (0); present (1). (Ostrom, 1976)
- number of sacral vertebrae: 5 or less (0), 6 or more (1). (adapted from Ostrom, 1976)
- fusion of all sacral vertebrae in adult: absent or partially fused (0); present, all completely fused (1).
- number of caudal vertebrae: 30 or more (0), 20-25 (1), or less than 15 (2).
- pygostyle: absent (0); present (1) (Sanz and Buscalioni, 1992)
- transition point on caudal vertebrae: distal, behind caudal 10 (0); proximal, no further back than caudal 10 (1) (Gauthier, 1986; Gatesy, 1996)
- middle and distal chevron shape: laterally compresses, expanded dorsoventrally (0); dorsoventrally flattened into a thin horizontal plate behind transition point (1). (adapted from Ostrom, 1976; Gauthier, 1986)
- hyposphene-hypantrum articulations on dorsal vertebrae: present (0); absent (1).
- pneumatic foramen on sacral vertebrae: present(0); absent (1).
- cervical neural spines: dorsoventrally tall, axially short (0); dorsoventrally short, axially elongate (1).
- length of middle and posterior caudal vertebrae: subequal to length of proximal caudals (0); elongate, at least 130% the length of the anterior caudals (1).
- prezygopophyses on middle and posterior caudal vertebrae: elongate, extend at least half the way across preceeding vertebral centrum (0); reduced or absent, extend over 25% or less the length of preceeding vertebral centrum (1).
- well-developed procoely in anterior caudal vertebrae: absent (0); present (1).
shoulder girdle
- scapula with tapered distal end: absent (0); present (1). (Ostrom, 1976)
- anterior projection of acromion process: absent (0); present, extends well anterior to glenoid fossa (1).
- scapulocoracoid articulation: immobile, sutural (0); mobile, cartilagenous (1) (adapted from Sanz et al., 1995)
- acroracoid of coracoid: absent, biceps tubercle small (0); absent, biceps tubercle large (1); present, extends above scapulocoracoid articulation, biceps tubercle large (1). (Perle et al., 1993; Walker, 1972; Ostrom, 1976)
- body of coracoid forming sharp angle with the body of the scapula: absent (0); present (1) (Ostrom, 1976; Tarsitano and Hecht, 1980)
- long axis of coracoid portion of glenoid relative to long axis of scapular portion of glenoid: alligned (0); form a 90 degree angle (1). (Chiappe)
- coracoid shape: short, rounded sternal border (0); elongate, subrectangular profile, rounded sternal border (1); elongate, strut-like (2). (adapted from Ostrom, 1976; Gauthier, 1986)
- dorsoventrally curved scapular shaft: absent (0); present (1).
- supracoracoid nerve foramen (incision) located in the medial margin of coracoid: absent (0); present (1). (Sanz et al., 1995)
- shape of fused clavicles: absent (0); present, interclavicular angle greater than 90 degrees (1); present, interclavicular angle less than 90 degrees (2). (Ostrom, 1976; Chiappe, 1991)
- ossified sternum: paired (0); fused or partially fused into a single structure (1); fused with a midline keel (2).
forelimb
- distal condyles of humerus located mainly on the distal (0), or cranial (1), aspect. (Perle et al., 1993)
- ventral tubercle of humerus projects distinctly caudally, separated from the humeral head by a deep capital incision: absent (0); present (1) (Sanz et al., 1995)
- well-developed transverse ligamental groove on the humerus: absent (0); present (1) (Sanz et al., 1995)
- humerus length relative to ulna: longer (0); subequal to or shorter (1). (Perle et al., 1993)
- dorsal condyle of ulna developed as a semilunar ridge: absent (0); present (1) (Sanz et al., 1995)
- ratio of diameter of shaft of radius to ulna: more than .70 (0); thinner than ulna, less than .70 (1). (adapted from Tarsitano and Hecht, 1980; Perle et al, 1993; Chiappe and Calvo, 1994)
- carpometacarpus: absent (0); present (1). (Sanz et al., 1995)
- extensor process on metacarpus: absent (0); present (1). (Sanz et al., 1995)
- metacarpal I greater than (0), or less than (1) one-third the length of MC II. (Tarsitano and Hecht, 1980; Gauthier, 1986)
- metacarpal III: straight (0); bowed laterally (1). (adapted from Tarsitano and Hecht, 1980; Gauthier, 1986)
- combined lengths III-1 and III-2 relative to III-3 on manus: longer (0); subequal (1); absent (2). (Gauthier, 1986)
- forelimb elongatation relative to presacral length: less than one half presacral length (0); 60-90% presacral length (1); more than 100% presacral length (2). (adapted from Ostrom, 1976; Gauthier, 1986)
- manus length relative to ulna length: manus shorter than or subequal to ulna (0); manus longer than ulna by at least 20% (1).
- proximal and distal ends of humerus: twisted, do not lie in the same plane (0); expanded into the same plane (1).
- ulnar distal condyle: transversely compressed and craniocaudally extended proximally in the same plane as the humero-ulnar flexion-extension movement (0); subtriangular in shape in distal view, with a dorsomedial condyle, and twisted more than 54 degrees with respect to the proximal end (1).
- shape of ulnar posterior margin: sigmoid (0); uniformly convex (1).
- distal radial carpal: proximodistally flattened (0); semilunate (1).
pelvic girdle
- length of preacetabular process of ilium relative to length of postabetabular process: subequal (0); twice as long (1). (adapted from Tarsitano and Hecht, 1980; Currie and Russell, 1988)
- postacetabular process depth: relatively deep, more than 50% depth a acetabulum (0); shallow, less than 50% depth at acetabulum, drawn back into a low, pointed process (1). (adapted from Ostrom, 1976; Novas, 1997)
- ratio of lengths of acetabulum to ilium: .15 or more (0); .11 to .13 (1); .10 or less (2). (adapted from Sanz et al., 1995)
- antitrochanter on ilium: absent or very small (0); present, prominent (1). (adapted from Ostrom, 1976; Perle et al., 1993)
- tubercle on dorsal margin of ilium above caudal acetabulum: absent (0); present (1).
- craniocaudal width of pubic peduncle on ilium relative to width of acetabulum: narrow, less than acetabular width (0); wide, exceeds acetabulum in width (1).
- pubic foot: projects cranially and caudally (0); projects caudally only (1); absent (2). (adapted from Ostrom, 1976; Gauthier, 1986; Sanz et al., 1995)
- angle of pubic shaft relative to the long axis of sacral vertebrae: projects cranially (0); subvertical (1); projects caudally (opisthopubic) (2).
- ratio of length of ischium to length of pubis: more than .66 (0), less than .66 (1). (adapted from Ostrom, 1976; Gauthier, 1986)
- contact between distal ischia: present (0); absent (1). (Tarsitano and Hecht, 1980; Sanz et al., 1995)
- obturator process on ischium: present, square (0); present, peaked and broadly triangular (1); rudimentary or absent (2).
- placement of obturator process: proximal (0); mid-shaft (1); distal (2).
- "postacetabular" process on proximal ischium behind iliac contact: absent (0); present, nearly contacts postacetabular process of ilium (1). (Ostrom, 1976)
- process on caudal border of midshaft of ischium: absent (0); present (1). (Ostrom, 1976)
- fusion of pelvic elements: absent (0); present (1). (Sanz et al., 1995)
- shape of ischial shaft: rod-like in part with a circular or subcircular cross section (0); mediolaterally compressed and plate-like along entire length (1).
- postacetabular blade on ilium: brevis shelf caudolaterally oriented, medial flange ventrally curved (0); postacetabular blade vertical, medial flange strongly reduced and perpendicular to iliac blade (1). (Novas, 1997)
- pubic apron: transversely broad, pubes fused for distal 2/3rds of their length (0); transversely narrowed, pubes fused only along the distal 1/3rd of their length (1).
- contact between distal pubes: present (0); absent (1).
- iliac fossa for m. cupedicus: present (0); absent (1).
hindlimb
- configuration of lesser trochanters: large and flange-like, separated from femoral shaft, head, and greater trochanter by a deep cleft (0); nearly confluent with the greater trochanter, and separated from the proximal head of femur by only a small cleft or groove (1); joined to greater trochanter to form an undivided trochanteric crest, proximal articular surface confluent with that of femoral head (2). (adapted from Ostrom, 1976; Tarsitano and Hecht, 1980; Gauthier, 1986; Currie and Russell, 1988; Perle et al., 1993)
- shape of fourth trochanter on femur: present, large (0); reduced to a low ridge or absent (1). (adapted from Tarsitano and Hecht, 1980; Gauthier, 1986; Chiappe, 1996)
- "posterior trochanter" on femur: absent (0); present, cranially placed (1); present, centered on the trochanteric crest (2). (adapted from Ostrom, 1986; Sanz et al., 1995)
- fossa for capital ligament: absent (0); present (1).
- popliteal fossa of femur bounded distally by transverse bridge: absent (0); present (1). (Perle et al., 1993)
- tibiofibular crest on the lateral condyle of distal femur: absent (0); present (1). (Sanz et al., 1995)
- femoral neck: present, constriction developed that separates trochanteric region from femoral head (0); absent (1) (adapted from Russell, 1969)
- proximal tibia: craniocaudal length twice that of mediolateral width (0); length and width of proximal tibia subequal (1).
- midshaft diameter of fibula relative to tibia: approximately one-fifth or more that of tibia (0); one-fifth or less that of tibia (1).
- orientation of iliofibularis tubercle on fibula: craniolateral (0); lateral (1); caudolateral or caudal (2). (Forster et al., 1996)
- deep fossa on medial side of proximal fibular head: present (0); absent (1).
- fibular articulation with the calcaneum: present (0); absent, fibula does not reach tarsus (1)
- fusion of proximal tarsals to crus: absent (0); present, partially fused (1); present, completely fused (2). (adapted from Ostrom, 1976; Forster et al., 1996)
- fusion of astragalus to calcaneum in adults: absent (0); present, partial to complete (1). (Currie and Peng, 1993)
- fusion of distal tarsals to metatarsus: absent (0); present (1). (Ostrom, 1976)
- fusion of metatarsus: absent or partially fused (0); completely fused (1). (Ostrom, 1976)
- metatarsal V: present (0); absent (1). (Ostrom, 1976)
- position of pes digit I: parallel to other digits (0); reversed to oppose other digits (1). (Ostrom, 1976)
- relative contributions of metatarsals II, III, and IV to ankle joint: all contribute approximately equally (0); partially or completely arctometatarsalian, MT III is nearly or completely eliminated from joint (1). (Currie and Peng, 1993)
- relative size of pes digit II: phalanges and ungual subequal in size and robustness to digits III and IV (0); developed into a robust, hyperextendable slashing digit with an enlarged sickle-like ungual (1). (Gauthier, 1986)
- ratio of length of tibia to length of femur: tibia no more than 15% longer than femur (0); tibia elongate, at least 25% longer than femur (1).
- extensor canal on tibiotarsus: absent (0); present (1).
- distal vascular forament on metatarsus (tarsometatarsus): absent (0); present (1).
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